1
IMPACT OF WATER DEFICIT ON LEAF AREA, LEAF AREA INDEX, SPECIFIC LEAF
2
WEIGHT AND YIELD OF BANANA CULTIVARS AND HYBRIDS
K. Krishna Surendar*1, D. Durga Devi1, I. Ravi2, P. Jeyakumar1 and K. Velayudham3
3
4
*1
5
2
3
6
7
Department of Crop Physiology, TNAU, Coimbatore-641 003
National Research Centre for Banana (ICAR), Thiruchirapalli.
Department of Farm Management, TNAU, Coimbatore-641 003
E-mail: [email protected]
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Abstract
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This study examined the relationship between the yield reduction by Leaf Area (LA), Leaf Area Index (LAI) and
10
Specific Leaf Weight (SLW). The field experiment was conducted at National Research Centre for Banana to
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screen the banana cultivars and hybrids for water deficit tolerance and to elucidate information on growth attribute
12
mechanism of banana cultivars and hybrids. Stress was imposed at different critical stages viz., 3rd, 5th, 7th and 9th
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month after planting. The stress was given by scheduling irrigation at the 50 per cent available soil moisture (ASM)
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characteristic during critical stages. The soil moisture content was analyzed by using pressure plate membrane
15
apparatus. In control plots, the irrigation was given at the ASM of 80 per cent with the soil water potential of around
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-6 bars and in the case of stressed plots; the irrigation was given when an ASM reached 50 per cent with the soil
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water potential of -14 bars. In stressed plots, 50 per cent ASM was reached around 30 days. In this present study
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conducted with twelve cultivars and hybrids with three replications. The data were analyzed by using split plot
19
design. The results revealed that the cultivars of Karpuravalli, Karpuravalli x Pisang Jajee, Saba, and
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Sannachenkathali recorded significantly higher yield (67.3, 52.4, 55.8 and 41.3 t/ha) and the magnitude of
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yield decrease was 12 per cent than the cultivars and hybrids of Matti, Pisang Jajee x Matti, Matti x
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Anaikomban and Anaikomban x Pisang Jajee (14.9, 11.1, 10.3 and 10.6 t/ha). Similarly, Karpuravalli,
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Karpuravalli x Pisang Jajee, Saba, and Sannachenkathali recorded significantly Leaf Area, Leaf Area Index and
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Specific Leaf Weight with lesser reduction percent were showed than the cultivars and hybrids of Matti,
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Pisang Jajee x Matti, Matti x Anaikomban and Anaikomban x Pisang Jajee.
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Key Words: Banana, Water deficit, Leaf Area, Leaf Area Index, Specific Leaf Weight and yield.
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Abbreviations: Leaf Area (LA), Leaf Area Index (LAI) and Specific Leaf Weight (SLW)
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(** - Highly significant) (* - significant)
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Introduction
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Soil water deficit is extremely damaging to the plant, in which can limit the production and productivity in
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crop plants (Santos and Carlesso, 1998), as well as provoking smaller growth during the vegetative period
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(Lobato et al., 2008), moreover it promotes flower abortion during the reproductive period (Pimentel,
33
2004). Banana is the ‘queen of tropical fruits’ and is one of the oldest fruits known to mankind from pre-
34
historic times. Today, it is the leading tropical fruit in the world market with a highly organized and
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developed industry. Water deficit is an major problem in banana growth and development. Although it
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has scores of definitions, it originates from a deficiency of precipitation over an extended period of time,
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usually a season or more. This deficiency results in a water shortage for some activity, group, or
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environmental sector. Banana plants respond and adopt to these stresses to survive under stress condition
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at the molecular and cellular levels as well as at the physiological and biochemical levels. Physiological
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responses to soil water deficit are the feature that is most likely to determine the response of the crop to
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irrigation. The banana plants are sensitivity to soil moisture stress is reflected in changes in reduced
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growth through reduced stomatal conductance and leaf size leads to reduction in photosynthetic pigments
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(Kallarackal et al., 1990) with increased leaf senescence (Turner, 1998). Leaf area is an important
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component that is closely related to the physiological processes controlling dry matter production and
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yield. Leaf area has been shown to influence the radiant energy interception, an important photosynthetic
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parameter in crop plants, showing positive relationship with net photosynthetic activity. Plants may
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respond to water deficit in different ways such as reducing leaf area, hence the transpiring surface (Meyer
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and Boyer, 1972). Leaf Area as one of the growth parameters also indicates the size of photosynthesizing
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apparatus. Leaf Area is a fundamental determinant of the total photosynthesis by the plant. Leaf Area
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showed a positive relationship with net photosynthetic activity. In banana, higher amount of LA on a
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shoot coincide with the emergence of the bunches (inflorescence) from the top of the pseudostem. After
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this, no new leaves are produced on that shoot because the bunch is terminal as the older leaves
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senescence (Turner, 1998). Turner (1998) found that water stress resulted in reduced LA leading to
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decreasing Leaf Area Index in banana. SLW is useful in understanding the means of the assimilates in leaf
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expansion. The SLA is a measure of LA per unit dry weight and it varies with cultivar, leaf position,
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growth stage and the environmental condition by Veerawirdh (1974). The SLW refers to photosynthetic
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efficiency and in turn higher total dry matter accumulation. It is the leaf dry weight per unit leaf area
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produced. Kramer (1983) found that water stress not only reduced LA but often increased leaf thickness,
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thereby increasing the weight per unit area i.e. in increased SLW. Thicker leaves aids in leaf water
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conservation because of the lower surface or lower volume ratio (Lopez et al., 1997). With this above
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background, the experiment aimed at evaluating the effects of the progressive water deficit, as well as to
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investigating the growth attributes behavior in twelve banana cultivars and hybrids submitted to water
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restriction during the different growth stages.
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Materials and Methods
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The experiment was carried out at National Research Centre for banana, Thiruchirapalli, during 2011-2012.
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The experiment consists of two treatments as considered as main plot and twelve cultivars and hybrids as taken as
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sub plots were laid out in split plot design with three replications. The main plots are, M1 (control) with the soil
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pressure maintained from -0.69 to -6.00 bar, M2 (water deficit) with the Soil pressure maintained from -0.69 to -
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14.00 bar. Soil pressure of -14.00 bar was reached at 30 days and measured by using soil moisture release curve and
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measured the soil moisture by using the pressure plate membrane apparatus. The sub plots are, S1: Karpuravalli
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(ABB), S2: Karpuravalli x Pisang Jajee, S3: Saba (ABB), S4: Sanna Chenkathali (AA), S5: Poovan (AAB), S6: Ney
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poovan (AB), S7: Anaikomban (AA), S8: Matti x Cultivar Rose, S9: Matti (AA), S10: Pisang Jajee x Matti, S11: Matti
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x Anaikomban and S12: Anaikomban x Pisang Jajee. The growth attributes of Leaf Area, Leaf Area Index and
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Specific Leaf Weight were measured during 3rd, 5th, 7th, 9th month after planting and at harvest stages of the crop.
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The procedure for measuring Leaf Area, Leaf Area Index and Specific Leaf Weight are given below:
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Leaf Area (LA)
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The leaf area was calculated by multiplying leaf length and breadth with the Constant factor 0.83
and number of green leaves and expressed in m2 (Hewitt, 1955).
Leaf Area = L x B x N x ‘K’
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L – Length of the leaf
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B – Breadth of the leaf
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N – Number of leaves
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‘K’ – Constant factor (0.83)
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85
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Leaf Area Index (LAI)
The Leaf Area Index (LAI) of functional leaves was calculated by employing the formula of
Williams (1946).
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Leaf area per plant
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LAI=
89
90
91
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Ground area occupied by the plant
Specific Leaf Weight (SLW)
The
Specific
Leaf
Weight
(SLW)
was
calculated
by
using
the
formula
of
Pearce et al. (1968) and expressed as mg cm-2.
93
94
---------------------------------------------
Leaf dry weight per plant (g)
SLW=
-----------------------------------------------------Leaf area per plant (m2)
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Results
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Leaf Area (LA)
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The Leaf Area was affected by water deficit in all the cultivars and hybrids as well as the interaction of M
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at S and S at M were significant (Table 1). Among the twelve cultivars and hybrids, Karpuravalli,
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Karpuravalli x Pisang Jajee, Saba, and Sannachenkathali had significant differences in leaf area under the
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irrigation at 50 per cent available soil moisture level. The highest Leaf Area was observed in Karpuravalli
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with very lesser reduction was noticed under the water deficit. The lowest Leaf Area was observed in
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Matti, Pisang Jajee x Matti, Matti x Anaikomban and Anaikomban x Pisang Jajee cultivars and hybrids
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under the water deficit, respectively. There was a high and positive correlation between Leaf Area and
105
yield water deficit conditions.
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Leaf Area Index (LAI)
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The result on LAI had similar effect were showed in all the growth stages and also all the cultivars and
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hybrids by water deficit. The interaction effects of M at S and S at M were significant differed at all the
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cultivars and hybrids (Table 2). Water deficit decreased LAI in banana cultivars and hybrids. Among the
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twelve cultivars and hybrids, Karpuravalli, Karpuravalli x Pisang Jajee, Saba, and Sannachenkathali had
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significant differences in LAI under the main plot treatments. The highest LAI were observed in
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Karpuravalli due to the water deficit. The lowest LAI was observed in Matti, Pisang Jajee x Matti, Matti x
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Anaikomban and Anaikomban x Pisang Jajee cultivars and hybrids under the water deficit, respectively.
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Specific Leaf Weight (SLW)
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The data on SLW was affected under water deficit as well as the interaction of M at S and S at M were
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significant at all stages of growth (Table 3). Water deficit reduced SLW in all the twelve banana cultivars
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and hybrids. Among the twelve cultivars and hybrids, Karpuravalli, Karpuravalli x Pisang Jajee, Saba, and
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Sannachenkathali had significant differences in SLW under the main plot treatments. The highest SLW
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was observed in Karpuravalli under the water deficit than the other cultivars and hybrids. The lowest SLW
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content was observed in Matti, Pisang Jajee x Matti, Matti x Anaikomban and Anaikomban x Pisang Jajee
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cultivars and hybrids under the water deficit, respectively. There was a high and positive correlation
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between SLW and yield water deficit conditions.
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Discussion
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Leaf area is a fundamental determinant of the total photosynthesis of a plant. Leaf area always shows a
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positive relationship with net photosynthetic activity, because leaf enlargement is attributed to increase in
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number and width of grana and also high degree of stacking of grana (Flore et al., 1985). Leaf area
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development is based on the length and width of leaf, in general, was very sensitive to water deficit in
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banana as reported by Turner (1981). The leaf length of banana reduced during water stress situation,
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which is associated with reduced organ development. Gardner et al. (1981) opined that water stress
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decreases the leaf area due to reduced cell division and cell enlargement which could be caused by
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accumulation of unexpanded cells during the cycle. According to the results obtained in the present study,
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the cultivars of Karpuravalli, Karpuravalli x Pisang jajee, Saba and Sannachenkathali showed a lesser
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reduction in leaf area in the range of 8 to 12 per cent due to water deficit over control. A 20 to 26 per cent
134
reduction in leaf area was registered by the cultivars of Poovan, Ney Poovan, Anaikomban and
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Anaikomban x Pisang jajee, whereas cultivars of Matti, Matti x Anaikomban, Matti x cultivar rose and
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Pisang jajee x Matti had higher reduction in leaf area of about 38 to 48 per cent over control. These results
137
were confirmed by the findings of Levy et al. (1978) observing that leaf area increases with an increase in
138
water supply because plants are able to photosynthesize more efficiently. This is because that an increased
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accumulation of photosynthates accelerates the pace of growth which in turn is reflected by vigorous plant
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growth. In banana, soil water regimes had a direct relationship on leaf width. There was an increase in leaf
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width with an increase in soil water regimes. This is because water is important for biochemical and
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physiological processes that lead to organ growth and development (Turner, 1972). A reduction in leaf
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area leading to reduced biomass accumulation and decreased growth and also leaf elongation of Kiwi fruit
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induced by water stress was a result of preferential partitioning of photosynthate to the roots and also
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shoots and thus affected leaf area development.
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Leaf Area Index (LAI) is one of the principle factors influencing canopy net photosynthesis of the crop
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plants (Hansen, 1982). The capacity of a canopy of leaves in a plantation to intercept light and fix carbon
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is measured by the LAI. Turner et al. (2007) reported that the optimum LAI for banana is 2 to 5. In banana
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plantation with LAI of 4.5 about 90% of the ground will be shaded at noon on a sunny day. This implies
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that about 90% of incoming radiation is being intercepted by the leaf canopy. Thus increasing LAI beyond
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this value is of little benefit to the plantation because most of the incoming solar radiation is already being
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intercepted (Turner et al., 2007). Drought stress induced changes in LAI, which duly reflected in biomass
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production (Kerby et al., 1990). Turner, (1998) found that water stress resulted in reduced LA leading to
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decreased LAI in banana. The lack of cell expansion due to water shortage would be determined by
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decreased LA rather than the number of leaves (Hsiao, 1973). In the present study also the effect of water
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deficit on LAI could be revealed. The cultivars like., Karpuravalli, Karpuravalli x Pisang jajee, Saba and
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Sannachenkathali showed a reduction of 8 to 12 per cent in LAI, whereas the cultivars like Poovan, Ney
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Poovan, Anaikomban and Anaikomban x Pisang jajee recorded 8 to 12 and 19 to 25 per cent reduction in
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LAI at 7th MAP over control. However, the other cultivars of Matti, Matti x Anaikomban, Matti x cultivar
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rose and Pisang jajee x Matti registered a higher reduction per cent of about 38 to 43 over control. As per
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the report of De Silva et al. (1979), reduction in LAI was observed due to acceleration of senescence
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under drought. According to Hoffman and Turner (1993), leaf growth rate was more sensitive to water
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stress.
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Specific Leaf Weight (SLW), a measure of thickness of leaf, has been reported to have a strong positive
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correlation with leaf photosynthesis in several crops as reported by Bowes et al.(1972). In many crop
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species, thicker leaves would have more number of mesophyll cells with high density of chlorophyll and,
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therefore, have a greater photosynthetic capacity than thinner leaves (Craufurd et al., 1999). Specific Leaf
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Weight is highly correlated with the development of reproductive organ namely flower and ultimately
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yield. As observed in the present study, Karpuravalli, Karpuravalli x Pisang jajee, Saba and
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Sannachenkathali recorded higher SLW with lesser reduction per cent of about 8 to 9 due to water deficit
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over control. The mechanism of maintaining higher SLW could be related to its thick leaves with more
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photosynthetic proteins per unit area of the leaf (Wells and Nugent, 1980). The higher reduction in SLW
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(24 to 26%) under stressed conditions in the cultivars of Matti, Matti x Anaikomban, Matti x cultivar rose
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and Pisang jajee x Matti could also be related to lesser number of mesophyll cells leads to lower
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photosynthetic efficiency (Gardner et al., 1985).
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Conclusion
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Plants respond to drought stress through alteration in physiological and biochemical processes. Our results
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showed that the growth attributes of Leaf Area, Leaf Area Index and Specific Leaf Weight decreased under the
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water deficit condition. The banana cultivars and hybrids of Karpuravalli, Karpuravalli x Pisang jajee,
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Saba and Sannachenkathali with lesser reduction in Leaf Area, Leaf Area Index and Specific Leaf Weight and
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also smaller bunch yield reduction when the plants endured water deficit. The findings of this research
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also showed that the Leaf Area, Leaf Area Index and Specific Leaf Weight can be used as a drought tolerance
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index to selection tolerant genotypes under water deficit conditions in banana cultivars and hybrids.
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Acknowledgment:
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The research have been supported and facilitated by National Research Centre for Banana (ICAR),
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Trichy. Tamil Nadu. India. I extend my sincere thanks to Dr. M. M. Mustaffa (Director) NRC for banana,
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Dr. D. Durga Devi (Professor) TNAU and Dr. I. Ravi (Sr. Scientist) NRC for banana for given proper
188
guidance during research.
189
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Table 1. Effect of water stress on leaf area (m2 plant-1) at different growth stages of banana cultivars and hybrids.
Treatments
3rd MAP
5th MAP
7th MAP
9th MAP
Harvest
Mean
M1
2.7
4.5
6.3
5.7
5.1
4.87
M2
2.1
3.9
5.1
4.5
3.9
3.89
Mean
2.41
4.20
5.66
5.14
4.48
4.38
SEd
0.023
0.037
0.054
0.053
0.039
CD (P= 0.05)
0.101
0.159
0.234
0.229
0.168
S1
5.1
8.8
13.7
12.0
9.3
9.77
S2
4.3
7.2
11.5
8.7
8.6
8.09
S3
3.8
7.1
7.9
7.9
7.1
6.76
S4
2.4
4.2
6.1
6.1
5.0
4.77
S5
2.6
4.6
6.5
5.8
5.2
4.92
S6
2.4
3.4
4.4
4.3
3.9
3.69
S7
2.0
3.1
3.9
3.7
3.4
3.22
S8
1.5
3.0
3.8
3.7
3.3
3.08
S9
1.5
2.8
2.8
2.7
1.9
2.34
S10
1.3
2.6
2.8
2.7
2.4
2.34
S11
1.1
2.2
2.3
2.0
1.9
1.88
S12
0.9
1.4
2.2
2.1
1.9
1.69
Mean
2.41
4.20
5.66
5.14
4.48
4.38
SEd
0.056
0.088
0.126
0.120
0.097
CD (P= 0.05)
0.114
0.179
0.254
0.243
0.197
Main plot
Sub plot
Interaction SEd
M at S
**
**
**
**
**
S at M
**
**
**
**
**
M at S
**
**
**
**
**
S at M
**
**
**
**
**
CD (P= 0.05)
251
252
253
Table 2. Effect of water stress on Leaf Area Index (LAI) at different growth stages of banana cultivars and hybrids.
Treatments
3rd MAP
5th MAP
7th MAP
9th MAP
Harvest
Mean
M1
0.69
1.13
1.57
1.43
1.27
1.22
M2
0.52
0.97
1.27
1.14
0.97
0.97
Mean
0.60
1.05
1.42
1.28
1.12
1.09
SEd
0.006
0.009
0.013
0.013
0.010
CD (P= 0.05)
0.027
0.042
0.057
0.056
0.046
S1
1.28
2.20
3.44
2.99
2.32
2.44
S2
1.08
1.81
2.88
2.18
2.15
2.02
S3
0.94
1.78
1.99
1.96
1.77
1.69
S4
0.60
1.06
1.53
1.52
1.25
1.19
S5
0.64
1.14
1.63
1.45
1.29
1.23
S6
0.60
0.86
1.11
1.08
0.97
0.92
S7
0.49
0.78
0.97
0.92
0.86
0.80
Main plot
Sub plot
S8
0.38
0.74
0.96
0.93
0.83
0.77
S9
0.38
0.70
0.70
0.67
0.49
0.59
S10
0.33
0.65
0.69
0.67
0.60
0.59
S11
0.28
0.54
0.57
0.51
0.46
0.47
S12
0.22
0.35
0.54
0.53
0.47
0.42
Mean
0.60
1.05
1.42
1.28
1.12
1.09
SEd
0.014
0.022
0.031
0.029
0.024
CD (P= 0.05)
0.028
0.045
0.064
0.060
0.049
M at S
**
**
**
**
**
S at M
**
**
**
**
**
M at S
**
**
**
**
**
S at M
**
**
**
**
**
Interaction SEd
CD (P= 0.05)
254
255
Table 3. Effect of water stress on Specific Leaf Weight (SLW: mg / cm2) at different growth stages of banana
256
cultivars and hybrids.
Treatments
3rd MAP
5th MAP
7th MAP
9th MAP
Harvest
Mean
M1
0.65
0.69
0.77
0.76
0.73
0.72
M2
0.53
0.57
0.65
0.64
0.61
0.60
Mean
0.59
0.63
0.71
0.70
0.67
0.66
SEd
0.008
0.005
0.007
0.007
0.007
CD (P= 0.05)
0.034
0.023
0.031
0.031
0.030
Main plot
Sub plot
S1
0.68
0.72
0.80
0.79
0.76
0.75
S2
0.67
0.71
0.79
0.78
0.75
0.74
S3
0.66
0.70
0.78
0.77
0.74
0.73
S4
0.65
0.69
0.77
0.76
0.73
0.72
S5
0.62
0.66
0.74
0.73
0.70
0.69
S6
0.60
0.64
0.72
0.71
0.68
0.67
S7
0.59
0.63
0.71
0.70
0.67
0.66
S8
0.56
0.60
0.68
0.67
0.64
0.63
S9
0.52
0.56
0.64
0.63
0.60
0.59
S10
0.51
0.55
0.63
0.62
0.59
0.58
S11
0.51
0.55
0.63
0.62
0.59
0.58
S12
0.50
0.54
0.62
0.61
0.58
0.57
Mean
0.59
0.63
0.71
0.70
0.67
0.66
SEd
0.007
0.008
0.009
0.009
0.008
CD (P= 0.05)
0.015
0.016
0.018
0.018
0.018
M at S
**
**
*
**
**
S at M
**
*
**
**
**
M at S
**
**
*
**
**
S at M
**
*
**
**
**
Interaction SEd
CD (P= 0.05)
257
258
259
260
261